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The sweet option: the importance of Aloe marlothii for opportunistic avian nectarivores PDF

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The sweet option: the importance of Aloe marlothii for opportunistic avian nectarivores Craig Symes L’option douce l’importance de Aloe marlothii pour des nectarivores aviens opportunistes. Les : especes de Aloe sont endemiques a l’Airique, oil elles sont repandues, offranr du nectar a une multitude d’especes animales. Cet article resume les resultats d une etude de trois ans, menee dans la Reserve naturelle du Suikerbosrand, Gauteng, Afrique du Sud, qui avait pour objectifd’examiner l’importance du nectar de A. marlothii pour les oiseaux. Le nectar est dilue et copieux et facilement accessible pour une multitude de nectarivores opportunistes (au moins 77 especes) qui agissent comme pollinisateurs legitimes. Summary. Aloe species are endemic and widespread in Africa, offering nectar to a host ofanimal species. This paper summarises the findings ofa three-year study at Suikerbosrand Nature Reserve, South Africa, investigating the importance ofA. marlothii nectar for birds. The nectar is dilute and copious and easily accessible to a host ofopportunistic neactarivores (at least 77 species) that act as legitimate pollinators. loe Linnaeus, species are widespread in the numbers of plants grow on rocky, north-facing Afrotropical region with a concentration slopes. During transects over six months almost of diversity in southern Africa (Holland 1978, half (38 of 83 species) of all species detected Reynolds 1969, Glen & Hardy 2000, Van Wyk were recorded feeding on nectar (Symes et al. & Smith 2005). A uniquely charismatic and 2008). An additional four species were recorded widespreadspeciesfound in SouthAfricansavannas feeding on nectar outside of transects (Symes et is Aloe marlothii A. Berger (Asphodelaceae), a al. 2008). In most observations, birds were seen m single-stemmed aloe that grows up to 6 in height probing the compact arrangement of flowers on (Reynolds 1969, Glen & Hardy 2000, Van Wyk the near-horizontal racemes. However, for many & Smith 2005). Flowers vary in colour throughout birds the bright wash of orange pollen dusted on their range from bright orange to red, and gaudy the facial area and belly was sufficient evidence for infloresencessuggestpollination bybirds (Reynolds a nectar ‘addiction’ (Fig. 1). Overall, throughout 1969, Glen & Hardy 2000, Van Wyk & Smith the range ofA. marlothii at least 77 bird species , 2005). The dilute nectar (c.12%), produced in are known to feed on the plant’s nectar. Although copious amounts (c.250 pl/flower) during the dry most of the species observed feeding on nectar winter period (June-September), when little else were residents, some species only appeared when is flowering, attracts a host ofopportunistic avian nectar was available during flowering. During this & nectarivores (Symes Nicolson 2008, Symes et period bird diversity and abundance increased al. 2008). Herein I summarise aspects of a three- significantly, suggesting that their arrival is year study investigating the symbiotic relationship dictated by the availability ofthe sugar-rich nectar ofA. marlothii and a suite of avian opportunistic of A. marlothii. One species that arrived in large nectarivores. numbers when A. marlothii flowered was Wattled Starling Creatophora cinerea. Most starlings An Aloe marlothii nectar oasis for birds (Sturnidae), mockingbirds (Mimidae), thrushes The first comprehensive documentation of birds (Turdidae) and robins (Muscicapidae) are unable feedingonA. marlothiinectarwas byOatley 964) to digest sucrose, a Cl2 sugar comprising two C6 (1 at Ndumu Game Reserve in northern KwaZulu- sugars, glucose and fructose (Martinez del Rio & Natal. His list included 43 nectar-feeding bird Stevens 1989, Martinez del Rio et al. 1992, Lotz & species belonging to 21 families. Although earlier Schondube 2006). However, most aloe nectars accounts exist (Marloth 1915), very little work has are glucose/lructose dominated (van Wyk et al. been done in understanding these animal-plant 1993) making their nectars suitable for a wider interactions. This prompted a study at a large range ofbird species. aloe ‘forest’ at Suikerbosrand Nature Reserve, Sunbird abundance during the flowering c.60 km south-east ofJohannesburg, where large period at Suikerbosrand was notably low and 178- BullABC Vol 17No 2 (2010) Importance ofAloe marlothiifor opportunisticavian nectarivores: Symes : ) could be explained by nectar characteristics. amounts of nectar ooze from the flower opening, Recent studies of plant—pollinator relationships many short-billed birds can access the nectar suggest that nectar concentration and volume are without damaging flowers. No birds have been important predictors of pollination syndromes, observed (like Cape White-eye Zosterops capensis with nectars of high volume (40-100 pl/flower) robbing nectar, confirming the legitimacy ofmost and low concentration (8-12% w/w) focused visitors as true pollinators. towards generalist pollinators, and nectars of low Of particular importance to the study was the volume (10-30 pl/flower) and high concentration use of stable isotopes, a technique that was in its (15-23% w/w) attracting specialist nectarivores infancy in the 1960s when Terry Oatley made & e.g., sunbirds (Johnson Nicolson 2008). his observations of opportunistic nectarivory at However, in the Eastern Cape the response of A. marlothii. Stable carbon isotopes are a useful birds to winter-flowering A. ferox might be quite tool in understanding animal diets because they different. Although the nectar in this ecologically quantify assimilated material, and depending on similar aloe is ofsimilar concentration and volume the tissue studied (which is related to turnover (180 pi, 12.5% w/w) it attracts large numbers rates of tissue) provide a window into different of Malachite Sunbird Nectarinia famosa so other dietary time periods. A. marlothii employs a factors (like a lack of major foodplants at this unique water-saving carbon fixation pathway season) besides nectar characteristics could be during photosynthesis (i.e. crassulacean acid CAM important determinants of plant visitors. metabolism or photosynthesis), which is The natural world is an incredible assemblage found in many plants of arid regions. Because of symbiotic relationships. Birds eat nectar but of this, it has a unique carbon isotopic signature how do aloes benefit? By feeding in a manner compared to C photosynthesising plants. By where pollen is deposited onto feathers, generalist measuring the carbon isotopic signature (SI3C) in birds act as important pollinators ofA. marlothii. whole blood before, during and after flowering, This was confirmed during two flowering seasons I was able to track the dietary shift of birds from by conducting pollinator exclusion experiments that of a C, photosynthetic source (i.e. broad- CAM (Symes et al. 2009). Honeybees Apis mellifera leaved plants) to nectar (with a isotope also visited aloes but in smaller numbers; they signature). Furthermore, by measuring carbon are probably excluded as pollinators because the isotope signatures ofCO, in breath samples was 1 nectar is more dilute than they prefer and because able to determine the importance of nectar sugars they were present in such low numbers at this as a readily available energy source for birds; this specific site; where numbers are greater their role is because carbon in breath represents immediately as pollinators might be more important. metabolised carbohydrates (Symes et al. in prep.). The tubular length of A. marlothii flowers The analysis of breath also indicated that analysis (c.33 mm) suggests a specialist bird-pollinated ofblood under-estimated the importance ofnectar syndrome (i.e. sunbirds). However, because large sugars for most nectar-feeding birds. Table 1. Numberofextantspecies in dominant nectarivore familiesforeach zoogeographic region (shaded), including sunbirds (Nectariniidae), sugarbirds (Promeropidae), honeyeaters (Meliphagidae) and hummingbirds (Trochilidae) (afterMaclean 1990). Tableau 1. Nombred’especes dans lesfamilies essentiellement nectarivores pourchaque region zoogeographique (hachure) souimangas (Nectariniidae), promerops (Promeropidae), meliphages (Meliphagidae) etcolibris (Trochilidae) (d'apres Maclean 1990), Neotropics Nearctic Palaearctic Afrotropics Indo-Malaya Australasia Total Trochilidae 324 13 0 0 0 0 337 Nectariniidae 0 0 2 78 39 2 121 Promeropidae 0 0 0 2 0 0 2 Meliphagidae 0 0 0 0 10 159 169 324 13 2 80 49 161 629 Importance ofAloe marlothiifor opportunisticavian nectarivores: Symes BullABC Vol 17No 2 (2010) - 179 Figure 1. African Red-eyed Bulbuls Pycnonotus nigricans Figure 2. Avian nectar feeders in Aloe marlothii forest feeding on Aloe marlothii flowers. The study site formed at Suikerbosrand Nature Reserve, Gauteng (a-f) and a region ofoverlap with Dark-capped Bulbul Pycnonotus common avian pollinators showing pollen dusted on facial tricolor (which were never observed, although hybrids area from feeding on nectar (g-j). were) (Craig Symes) (a) Immature male Malachite Sunbird Nectariniafamosa Bulbuls brunoirs Pycnonotus nigricans se nourrissant sur les and African Red-eyed Bulbul Pycnonotus nigricans; (b) fleurs deAloe marlothii. Le site d’etude formait une zone Aloe marlothii inflorescences; (c) Cape White-eye Zosterops de chevauchement avec le Bulbul des jardins Pycnonotus capensis; (d) immature male Malachite Sunbird Nectarinia barbatus tricolor (qui n’a jamais ete observe, bien que des famosa probing flowers on raceme; (e) Wattled Starling hybrides ont ete vus) (Craig Symes) Creatophora cinerea\ (f) inflorescence with Fiscal Flycatcher Sigelussilens perched on open flowers; (g) adult male Cape Weaver Ploceus capensis; (h) Black-collared Barbet Lybius torquatus; (i) female Wattled Starling Creatophora cinerea; Red-faced Mousebird Urocolius indicus (Craig Symes) (j) Oiseaux se nourrissant de nectar dans une foret deAloe marlothii, Reserve naturelle du Suikerbosrand, Gauteng, Afrique du Sud (a-f) et pollinisateurs aviens communs OK avec du pollen sur leur face apres avoir pris du nectar (g-j)- (a) Souimanga malachite Nectariniafamosa male immature et Bulbul brunoir Pycnonotus nigricans (b) inflorescences ; deAloe marlothii (c) Zosterops du Cap Zosterops capensis ; ; (d) Souimanga malachite male immature explorant des fleurs (e) Etourneau caroncule Creatophora cinerea (f) ; ; inflorescence avec un Gobemouche fiscal Sigelussilens perche sur des fleurs ouvertes ; (g) Tisserin du Cap Ploceus capensis male adulte (h) Barbican a collier Lybius ; torquatus (i) Etourneau caroncule Creatophora cinerea ; femelle Coliou quiriva Urocolius indicus (Craig Symes) ; (j) 180- BullABC Vol 17No 2 (2010) Importance ofAloe marlothiifor opportunisticavian nectarivores: Symes Importance ofAloe marlothiifor opportunisticavian nectarivores: Symes BullABC Vol 17No 2 (2010)-181 , Avian nectarivory in the Afrotropics of opportunistic nectar feeders for a single aloe Avian nectarivory is most broadly dominated by species in southern Africa. Therefore, for the rest three families: the New World hummingbirds ofthe continent this list could be greatly increased. (Trochilidae), the Old World sunbirds To date the diversity ofopportunistic nectarivores (Nectariniidae)andtheAustralo-Pacifichoneyeaters appears greater than in other zoogeographic (Meliphagidae) (Maclean 1990; Table 1). The regions, and is probably explained by the different sugarbirds (Promeropidae, two species) were once feeding roles of birds in different regions. In considered a unique nectar-feeding family of the Australo-Pacific region the equivalent role Africa, although arecentstudysuggestsadditions to of opportunistic nectarivores might be filled by this taxon (i.e. Grey-chested Illadopsis Kakamega honeyeaters with broader dietary requirements, poliothorax Spot-throat Modulatrix stictigula whilst in the Afrotropics the role ofopportunistic and Dapple-throat Arcanator orostruthus that nectarivores may be filled by several families ) are not nectar feeders (Beresford et al. 2005). that do not typically feed on nectar (Keast Nectar-feeding is not confined to specialist taxa 1985; Appendix). Furthermore, competition and occurs in a wide range of other families. of opportunistic nectarivores with specialist In the Neotropics the number of nectar-feeding nectarivores (i.e. sunbirds) in theAfrotropics could specialists, besides hummingbirds, is comparatively be high, thereby explaining the low number of low and includes members of the Dacnini and sunbird species in the Afrotropics compared to the Coerebini, tribes ofthe Thraupidae and Parulidae number of hummingbird and honeyeater species respectively (Burns et al. 2003, Dickinson 2003). in the Neotropics and Australasia respectively, In some regions where these major nectarivore where there are fewer opportunistic nectarivores families have not colonised, and monopolised (Table 1). An alternative explanation may relate the nectar-feeding niche, diversification within to degrees ofweather predictability and associated & other families has occurred (Nicolson Fleming plant assemblages and flowering patterns in each 2003). These include, for example, the Hawaiian of the regions. honeycreepers (Drepanidinae) (Pratt 2005), the asities and sunbird-asities (Philepittidae; four Other interactions involving aloes species) of Madagascar (Prum 1993, Irestedt Aloe marlothii nectar is important for many other et al. 2001), the Irenidae (fairy-bluebirds; two organisms besides birds and the use of nectar species), Chloropseidae (leafbirds; eight species) might be an important determinant of local and Aegithinidae (ioras; four species) of the Indo- movements for Chacma Baboons Papio hamadryas Malayan Region (Wells et al. 2003a, Dickinson ursinus. Observations suggest that troops with 2003, Fuchs et al. 2006), and the Dicaeidae ranges that overlap with A. marlothii make greater (flowerpeckers, 42 species) ofAustralasia (Delacour use of the aloe forest during flowering, utilising & 1944, Beecher 1953, Sibley Ahlquist, 1991, nectar and succulent leaves as a food and water Ericson & Johansson 2003). The enigmatic source (pers. obs.). During periods of low rainfall Lorinae (Psittacidae) of Australasia are also other mammal species might make use of Aloe uniquely adapted for nectarivory with a brush-tip nectar but may occur in such low numbers that tongue (Forshaw 2006), as is a single member of the scale ofcompetition with birds is low. the Timalidae (babblers), the Fire-tailed Myzornis Aloe marlothii is also important to birds for Myzornispyrrhoura (Wells etal. 2003b). However, many other reasons. The most abundant bird the Melanocharitidae (berrypeckers, 12 species) species recorded during censuses was Laughing of New Guinea lack the specialised structure Dove Streptopelia senegalensis, a species that of the tongue for feeding on nectar (Beehler seldom (if ever) fed on nectar. It was observed et al. 1986). The stitchbird (Notiomystidae), nesting during drier months (August-September) recently relegated to a monotypic family (from on horizontal leaves of tall aloes. Although the Meliphagidae), is a nectar-feeder endemic to New spines of aloes afford protection from predators, Zealand (Driskel et al. 2007). baboons are able to ascend plants to access nectar In the Afrotropics the overall diversity of or rob nests. Also, avian predators might still pose opportunistic nectar-feeding birds is unknown. a threat; in one instance a Pied Crow Corpus alhus This study has highlighted the high diversity was recorded robbing a nest with eggs (pers. obs.); 182-BullABC Vol 17 No 2 (2010) Importance ofAloe marlothiifor opportunistic avian nectarivores: Symes the latter species was, however, never observed Driskel, A., Christidis, L., Gill, B. J., Boles, W. E., feeding on nectar. Aloes also benefited birds Barker, F. K. & Longmore, N. W. 2007. A new in other ways besides providing nectar. Water endemic family of New Zealand passerine birds: collected after rains in the horizontal leaves was adding heat to the biodiversity hotspot. Austral. J. used as a drinking and bathing source by birds. Zool. 55: 73-78. Barbets excavated nesting cavities in trunks of Ericson, P. G. P. & Johansson, U. S. 2003. Phylogeny tall aloes, and the dry leaf skirt was used as a ofPasserida (Aves: Passeriformes) based on nuclear & protected site for birds to construct nests (e.g. and mitochondrial sequence data. Mol. Phylo. Fiscal Flycatcher Sigelus silens). Evol. 29: 126-138. Forshaw, M. 2006. Parrots of the World: An J. Conclusion Identification Guide. Princeton, NJ: Princeton The list of birds recorded feeding on aloes has FuchUsn,iversitFyjePlrdessas,. & Pasquet, E. 2006. An increased from that previously known and includes J., J. ancient African radiation of corvoid birds (Aves: at least 101 species in 26 families (Appendix). Passeriformes) detected by mitochondrial and Some species might irregularly feed on nectar nuclear sequence data. Zool. Scripta 35: 375-385. and observations of them doing so might be rare: Glen, H. F. & Hardy, D. S. 2000. Fascicle 1: Aloaceae A. marlothii is a widespread species in southern (First part): Aloe. In Germishuizen, G. (ed.) Flora Africa and the number of species feeding on its ofSouthernAfrica. Vol. 5. Part 1. Pretoria: National nectar could therefore exceed 77 species. This Botanical Institute. phenomenal diversityofopportunistic nectarivores Hockey, P. A. R., Dean, W. R. & Ryan, P. G. (eds.) J. is possibly the highest number of bird species 2005. Roberts' Birds of Southern Africa. Seventh recorded feeding on the nectar of a single plant edn. Cape Town: Trustees of the John Voelcker species worldwide (Appendix 1). Bird Book Fund. Hoffman, M. T. 1988. The pollination ecology ofAloe Acknowledgements ferox Mill. S. Afr. J. Bot. 54: 345-350. This study was funded by the National Research Holland, P. G. 1978. An evolutionary biogeography of Foundation (South Africa). Darren Pietersen and the genus Aloe.J. Biogeogr. 5: 213-226. & Tracy Symes are thanked for assistance with the field Irestedt, M.,Johansson, U. S., Parsons, T. J. Ericson, work. Franqoise Dowsett-Lemaire commented on the P. G. P. 2001. Phylogeny of major lineages of submitted manuscript. suboscines (Passeriformes) analysed by nuclear DNA sequence data Avian Biol. 32: 15-25. .J. References Johnson, S. D. & Nicolson, S. W. 2008. Evolutionary Beecher, W. 1953. A phylogeny ofoscines. Auk 70: associationsbetweennectarpropertiesandspecificity J. 270-333. in bird pollination systems. Biol. Lett. 4: 49-52. Beehler, B. M., Pratt, T. K. & Zimmerman, D. Keast,A. 1985.An introductoryecologicalbiogeography A. 1986. Birds of New Guinea. Princeton, NJ: ofthe Australo-Pacific Meliphagidae. New Zealand Zool. 12: 605-622. Princeton University Press. J. & Beresford, P., Barker, F. K., Ryan, P. G. & Crowe, Lotz, C. N. Schondube, J. E. 2005 Sugar preferences T. M. 2005. African endemics span the tree of in nectar- and fruit-eatingbirds: behavioral patterns and physiological causes. Biotropica 38: 3-15. songbirds (Passeri): molecular systematics ofseveral Maclean, G. L. 1990. Ornithology for Africa. evolutionary 'enigmas’. Proc. Roy. Soc. 272(1565): Pietermaritzburg: University ofNatal Press. 849-858. Burns, K. J., Hackett, S. J. & Klein, N. K. 2003. MaclSeiaxnt,h Ge.dnL.. 1C9a9p3e. RToboewrnts:’ BTirrudsstoefesSouotfhetrhneAfJriochan. Phylogenetic relationships of Neotropical Voelcker Bird Book Fund. honeycreepers and the evolution of feeding morphology./. Avian Biol. 34: 360-370. Marloth, R. 1915. The Flora of South Africa, With Delacour,J. 1944. Arevision ofthefamilyNectariniidae SPlpaenctisa.l VSoynlo.pt4i.caClaTpaebleTsowonf:theDarGteenreraBrootfheHrisghe&r (sunbirds). Zoologica 29: 17-38. Co. Dickinson, E. C. (ed.) 2003. The Howard and Moore & Martinez del Rio, C. Stevens, B. R. 1989. Complete Checklist ofthe Birds ofthe World. Third Physiological constraint on feeding behavior: edn. Princeton, NJ: Princeton University Press. Importance ofAloe marlothiifor opportunisticavian nectarivoies. Symes BullABC Vol 17No 2 (2010) -183 — . intestinal membrane disaccharidases oi the starling. floweringofAloemarlothii'. a nectaroasis duringdry Science 243: 794-796. South African winters./ Ornithol. 149: 13-22. Martinez del Rio, C., Baker, H. G. & Baker, I. Symes, C. T., Human, H. & Nicolson, S. W. 2009. 1992. Ecological and evolutionary implications Appearances can be deceiving: pollination in two of digestive processes: bird preferences and the sympatric winter-flowering Aloe species. S. Afr. J. sugar constituents of floral nectar and fruit pulp. Bot. doi: 10.10l6/j.sajb.2009.08.008. Experientia 48: 344-551. Symes, C. T., McKechnie, A. E., Nicolson, S. W. Nicolson, S. W. & Fleming, P. A. 2003. Nectar as & Woodborne, S. M. in prep. The nutritional food for birds: the physiological consequences of significance of winter-flowering Aloe marlothii for drinking dilute sugar solutions. Plant Syst. Evol. opportunistic avian nectarivores. & 238: 139-153. Van Wyk, B.-E. Smith, G. 2005. Guide to theAloes Oatley, T. B. 1964. The probing of aloe flowers by ofSouthernAfrica. Pretoria: Briza Publications. birds. Lammergeyer 3: 2-8. Van Wyk, B.-E., Whitehead, C. S„ Glen, H. F„ — & Oatley,T. B. 2001. Gardenvisitors sunbirds.H/rzAz Hardy, D. S., Van Jaarsveld, E. J. Smith, G. F. & Birds Birding6(3): 26-31 1993. Nectar sugar composition in the subfamily Oatley, T. B. & Skead, D. M. 1972. Nectar feeding by Alooideae (Asphodelaceae). Biochem. Syst. Ecol. 21: South African birds. Lammergeyer 15: 68-69. 249-253. Pratt, H. D. 2005. The Hawaiian Honeycreepers. Wells, D. R., Dickinson, E. C. & Dekker, R. W. R. Oxford: Oxford University Press. 2003a. Systematic notes on Asian birds. 36. J. Prum, R. O. 1993. Phylogeny, biogeography, and A preliminary review of the Chloropseidae and evolution of the broadbills (Eurylaimidae) and Iredinae. Zool. Verhand. (Leiden) 344: 25-42. asities (Philepittidae) based on morphology. Auk Wells, D. R., Dickinson, E. C. & Dekker, R. W. R. 110: 304-324. 2003b. Systematic notes on Asian birds. 34. J. Reynolds, G. W. 1969. TheAloes ofSouthAfrica. Cape A preliminary review of the Aegithinidae. Zool. Town: A. A. Balkema. Verhand. (Leiden) 344: 7-15. Skead, C. 1967. The Sunbirds ofSouthern Africa. J. School ofAnimal, Plant and Environmental Sciences, Cape Town: A. A. Balkema. Symes, C. T. & Nicolson, S. W. 2008. Production of University ofthe Witwatersrand, P/Bag3, WITS, 2050, SouthAfrica. E-mail: [email protected] copious dilute nectar in the bird-pollinated African succulent Aloe marlothii (Asphodelaceae). S. Afr. J. Received 3 December 2008; revision accepted 21 Bot. 74: 598-605. September 2009. Symes, C. T., Nicolson, S. W. & McKechnie, A. E. 2008. Response of avian nectarivores to the Appendix 1. Birdspeciesrecordedfeeding on,orsuspectedoffeedingon,Aloemarlothiinectar. Dominantfeedingguild(afterMaclean 1993):fr=frugivore,gr=granivore, in=insectivore, ne=nectarivore,om =omnivore. Levelofnectarivory(afterOatley 1964): 0=non-feederbutpresentthereforesuspectedfeeder, 1 =occasional, 2=casual,3=regular,4=addict, 5=truenectarivore. SNR=recordedatSuikerbosrand NatureReserve. References: 1 =Oatley(1964), 2=Skead (1967), 3=Oatley&Skead (1972),4=Hoffman (1988)(recordsattributedtoA. ferox), 5=Maclean (1993),6=Oatley(2001), 7=pers. obs„8=thisstudy, 9=A. Craig pers.comm, (recordsattributedtoA. ferox), 10= B.deBoerpers. comm., 11 = M. Kriekpers.comm., 12=C. Botesunpubl. data. Taxonomyfollows Hockeyeta/.(2005). *Buphaguserythrorhynchuswasobservedforthefirsttimeatthestudysite(firstknownrecordforSNR)onlyduringtheflowering periodandisbyassociationwithA. marlothiirecordedasa nectarfeeder. Annexe 1. Especesd'oiseauxayanteteobserveesentraindesenourrirdu nectardeAloemarlothiioususpecteesdelefaire. Regimealimentairedominant(d’apres Maclean 1993) :fr=frugivore,gr=granivore, in=insectivore, ne=nectarivore,om=omnivore. Niveaudenectivorie(d’apresOatley 1964) :0= n’a paseteobservesenourrissantdenectarmaispresentetainsisuspectedelefaire, 1 =accidentel,2=occasionnel, 3 =regulier,4=tresregulier, 5=vrainectarivore. SNR=observedansla ReservenaturelleduSuikerbosrand. References : 1 =Oatley(1964),2=Skead(1967),3=Oatley&Skead (1972),4= Hoffman (1988)(mentionsattribueesaA. ferox),5=Maclean(1993),6=Oatley (2001),7=obs. pers., 8=cetteetude,9=A.Craigcomm. pers. (mentionsattribueesaA. ferox), 10=B.de Boercomm, pers., 11 =M. Kriekcomm, pers., 12=C. Botes donneesnon publiees. LataxonomiesuitHockeyetal. (2005). *Buphaguserythrorhynchusaeteobservepourla premierefoissurlesited’etude(premierementionconnuepourSNR)uniquementpendantla periodedefloraisonet estconsiderecommeun nectarivoreparsonassociationavecA. marlothii. 184 - BullABC Vol 17No 2 (2010) Importance ofAloe marlothiifor opportunistic avian nectarivores: Symes 3 Speciesandfamily Scientific Name Feeding Guild Level ofnectarivory SNR Refs. Indicatoridae LesserHoneyguide Indicatorminor In 1 1 3 Picidae CardinalWoodpecker Dendropicosfuscescens In 1 1 8 Lybiidae White-eared Barbel Stactolaemaleucotis Fr/ln 2 1 Yellow-rumpedTinkerbird Pogoniulusbilineatus Fr/ln 1 - 3 Yellow-frontedTinkerbird Pogoniuluschrysoconus Fr/ln 2 - 10 Red-frontedTinkerbird Pogoniuluspusillus Fr/ln 2 - 1 AcaciaPied Barbet Tricholaemaleucomelas Fr 2 ' 1 1,3,8 Black-collared Barbet Lybiustorquatus Fr/ln 2 1 1,3,8 Crested Barbet Trachyphonusvaillantii In/Fr 1 1 3,8 Phoeniculidae GreenWoodhoopoe Phoeniculuspurpureus In 1 1 3 Rhinopomastidae CommonScimitarbill Rhinopomastuscyanomelas In 1 1 1,3,8,11 Coliidae White-backed Mousebird Coliuscolius Fr/Fo 2 1 3 Speckled Mousebird Coliusstriatus Fr/Fo 4 1 1,2,3,8,12 Red-faced Mousebird Uroco/iusindicus Fr/Fo 4 1 1,3,8 Psittacidae Brown-headed Parrot Poicephaluscryploxanthus Fr 4 - 1 Musophagidae GreyGo-away-bird Corythaixoidesconcolor Fr/Fo 3 1 3,7 Columbidae Laughing Dove Slreptopeliasenegalensis Gr/ln 1 1 1,3,8 CapeTurtleDove Streptopeliacapicola Gr/ln 1 1 1,3 Oriolidae Black-headed Oriole Orioluslarvatus In/Fr 3 1 1,2,3,9,10 Dicruridae Fork-tailed Drongo Dicrurusadsimilis In/Ca 3 - 1,2,3,9,12 Malaconotidae Brown-crownedTchagra Tchagraaustralis In 1 1 8 Southern Boubou Laniariusferrugineus In/Fr 3 1 1,3 Corvidae Pied Crow Cornsa/bus Ca/Gr/Fr 1 1 3 White-necked Raven Cornsalbicollis Ca 3 1 Garlandin 1. Paridae GreyPendulineTit Anthoscopuscaroli In 3 - 1 Southern BlackTit Parusniger In 2 1,3 Pycnonotidae Dark-capped Bulbul Pycnonotustricolor Fr/ln 4 1 1,3,11 African Red-eyed Bulbul Pycnonotusnigricans Fr/ln 4 1 3,8 Cape Bulbul Pycnonotuscapensis Fr 3 - 5 SombreGreenbul Andropadusimportunus In/Fr 4 - 1,3,12 Terrestrial Brownbul Phyllastrephusterrestris In/Fr 4 1 Sylviidae CapeGrassbird Sphenoeacusafar In 1 1 3,8 Yellow-bellied Eremomela Eremomelaicteropygialis In 1 1 3 Burnt-necked Eremomela Eremomelausticollis In 2 - 1 Long-billed Crombec Sylviettarufescens In 2 1 1,3,8 Arrow-marked Babbler Turdoidesjardineii In/Fr 2 1,3 Importance ofAloe marlothiifor opportunisticavian nectarivores: Symes BullABC Vol 17No 2 (2010) -185 9 Speciesandfamily Scientific Name FeedingGuild Levelofnectarivory SNR Refs. Chestnut-ventedTit-Babbler Parisomasubcaeruleum In/Fr 1 1 3,8 Zosteropidae AfricanYellowWhite-eye Zosteropssenegalensis In/Fr 3 _ 1 CapeWhite-eye Zosteropscapensis In/Fr 4 1 2,3,8,9,10,12 Cisticolidae Rattling Cisticola Cisticolachiniana In 4 1 1,3,8 Neddicky Cisticolafulvicapilla In 4 1 1,3,8 Tawny-flanked Prinia Priniasubflava In 3 1 1,3 Black-chested Prinia Priniaflavicans In 3 1 3,8 KarooPrinia Priniamaculosa In 1 - 3 Bar-throatedApalis Apalisthoracica In 2 1 8 Yellow-breastedApalis Apalisflavida In/Fr 1 - 1,3 Rudd’sApalis Apalisruddi In 3 - 3 Muscicapidae 1 CapeRockThrush Monticolarupestris In/Fr 1 1 3,8,12 PaleFlycatcher Bradornispallidus In/Fr 2 - 1 Southern BlackFlycatcher Melaenornispammelaina In/Fr 2 - 1 FiscalFlycatcher Sigelussilens In/Fr 3 1 1,3,8 African DuskyFlycatcher Muscicapaadusta In/Fr 2 - 12 Cape Robin-Chat Cossyphacaffra In/Fr 1 1 8 White-browedScrub-Robin Cercotrichasleucophrys In/Fr 2 - 1 Buff-streakedChat Oenanthebifasciata In 3 - 3 Mocking CliffChat Thamnolaeacinnamomeiventris In/Fr 1 1 3 Sturnidae Red-wingedStarling Onychognathusmorio Fr/ln 3 1 3,4,7,9,12 Black-belliedStarling Lamprotorniscorruscus Fr/ln 3 - 1,3 CapeGlossyStarling Lamprotornisnitens In/Fr 3 1 1,3,7,8,9 Pied Starling Spreobicolor In/Fr 3 1 3,9 Wattled Starling Creatophoracinerea In/Fr 1 1 8 Red-billedOxpecker* Buphaguserythrorhynchus In 0 1 8 Nectarinidae OliveSunbird Cyanomitraolivacea Hello 5 - - mom GreySunbird Cyanomitra Hello 5 - - AmethystSunbird Chalcomitraamethystina Ne/ln 5 1 2,4,7,9,12 Scarlet-chestedSunbird Chalcomitrasenegalensis Ne/ln 5 - 2,7 BronzySunbird Nectariniakilimensis Ne/ln 5 - 2 MalachiteSunbird Nectariniafamosa Ne/ln 5 1 2,7,8,9,12 Collared Sunbird Hedydipnacollaris Ne/ln 5 - 7,12 Southern Double-collaredSunbird Cinnyrischalybeus Ne/ln 5 - 2,4,7,9 GreaterDouble-collared Sunbird Cinnyrisafer Ne/ln 5 - 2,7,9,12 White-belliedSunbird Cinnyristalatala Ne/ln 5 1 2,7,8 DuskySunbird Cinnyrisfuscus Ne/ln 5 - 2,6,9 MaricoSunbird Cinnyrismariquensis Ne/ln 5 - 2,6 Promeropidae Gurney'sSugarbird Promeropsgurneyi Ne/ln 5 _ 2 CapeSugarbird Promeropscater Ne/ln 5 2 Ploceidae LesserMaskedWeaver Ploceusintermedius In/Gr 3 - 5 SpectacledWeaver Ploceusocularis In/Fr/Gr 4 - 1 CapeWeaver Ploceuscapensis In/Gr 3 1 2,3,4, 7,8, U YellowWeaver Ploceussubaureus In/Gr/Fr 4 - Southern MaskedWeaver Ploceusvelatus In/Gr 4 1 1,3,8 186-BullABC Vol 17No 2 (2010) Importance ofAloe marlothiiforopportunisticavian nectarivores: Synies Species andfamily Scientific Name Feeding Guild Level ofnectarivory SNR Refs. VillageWeaver Ploceuscucullatus In/Gr 4 1 2,3 , Dark-backedWeaver Ploceusbicolor In/Fr 2 1 White-wingedWidowbird Euplectesalbonotatus Gr/ln 1 3 Red-collaredWidowbird Euplectesardens Gr/ln 1 1 3,8 Estrildidae Black-facedWaxbill Estrildaerythronotos Gr/ln 1 1 3,8 CommonWaxbill Estrildaastrild Gr/Fr 1 1 3 Violet-earedWaxbill Granatinagranatina Gr 1 1 8 Green-winged Pytilia Pytiliamelba Gr 1 1 8 Jameson’sFirefinch Lagonostictarhodopareia Gr 1 1 8 Passeridae HouseSparrow Passerdomesticus Gr/Fr/ln 1 1 3 CapeSparrow Passermelanurus Gr/ln/Fr 1 1 3 SouthernGrey-headed Sparrow Passerdiffusus Gr/ln 2 1 3,8 Yellow-throated Petronia Petroniasuperciliaris In/Gr 2 1 1,2,3 Fringillidae Black-throated Canary Seriousatrogularis Gr/ln 1 1 3,8 Yellow-frontedCanary Seriousmozambicus Gr/ln/Fo 3 1 1,2,3 YellowCanary Seriousflaviventris Gr/ln/Fo 2 1 3 BrimstoneCanary Serioussuiphuratus Gr/Fr/Fo 3 1 1,2,3 Streaky-headed Seedeater Seriousgularis Gr/Fo 3 1 1,3,4,8,12 Cinnamon-breasted Bunting Emberizatahapisi Gr/ln 1 1 8 Cape Bunting Emberizacapensis Gr 1 1 8 Total numberofspecies 101 Join our great value tours to Madagascar, Namibia, Cameroon, Malawi & more. Book Cape Town Pelagics & day trips. Ask about last-minute discounts. Birding Cape Town Contact Callan Cohen Pelagics Call +27 21 531 9148 www.birdingafrica.com Email [email protected] www.capetownpelagics.com Importance ofAloe marlothiifor opportunisticavian nectarivoies: Symes BullABC Vol 17No 2 (2010) -187

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